Eur J Appl Physiol (2000) 83: 566±572 Ó Springer-Verlag 2000

ORIGINAL ARTICLE

Johan Andersson á Erika Schagatay

Anna GisleÂn á Boris Holm

Cardiovascular responses to cold-water immersions of the forearm

and face, and their relationship to apnoea

Accepted: 18 July 2000

Abstract Apnoea as well as cold stimulation of the face

or the extremities elicits marked cardiovascular re¯exes
Introduction
in humans. The purpose of this study was to investigate
Apnoea or cold stimulation of either the face or the
whether forearm immersion in cold water has any e€ect
extremities elicits marked cardiovascular re¯exes in hu-
on the cardiovascular responses to face immersion and
mans. Stimulation of both the sympathetic and para-
apnoea. We recorded cardiovascular responses to cold-
sympathetic nervous systems are involved in evoking
water immersions of the forearm and face in 19 (part I)
these re¯exes. The human diving response is initiated by
and 23 subjects (part II). The experimental protocol was
apnoea and is augmented by immersion of the face in
divided in two parts, each part containing four tests: I1,
cold water (Gooden 1994). It is characterised by a
forearm immersion during eupnoea; I2, face immersion
reduction in heart rate, which is caused by an increased
during eupnoea; I3, forearm and face immersion during
parasympathetic activity via the vagus nerve that
eupnoea; I4, face immersion during apnoea; II1, apnoea
inhibits the cardiac pacemaker, and an alpha-adrenergic
without immersion; II2, forearm immersion during
sympathetic vasoconstriction in selective vascular beds
apnoea; II3, face immersion during apnoea; and II4,
(Heistad et al. 1968; Frey et al. 1980a; Gooden 1994).
forearm and face immersion during apnoea. The water
The reduction in cardiac output is not balanced to the
temperature was 9±11 °C. Cold-water immersion of
vasoconstriction and, as a consequence, blood pressure
either the forearm or face was enough to elicit the most
is increased. The diving response is augmented by
pronounced thermoregulatory vasoconstriction during
stimulation of cold-receptors in the upper facial region,
both eupnoea and apnoea. During eupnoea, heart rate
which is innervated by the ophthalmic branch of the
responses to forearm immersion (3% increase) and face
trigeminal nerve (Schuitema and Holm 1988).
immersion (9% decrease) were additive during concur-
Cold stimulation of the extremities during eupnoea
rent stimulation (3% decrease). During apnoea, the
(the cold pressure test) elicits an inhibition of the para-
heart rate responses were not a€ected by the forearm
sympathetic activity and increases the beta-adrenergic
immersion. The oxygen-conserving diving response
sympathetic activity to the heart, resulting in a tachy-
seems to dominate over thermoregulatory responses in
cardia. At the same time, an alpha-adrenergic sympa-
the threat of asphyxia. During breathing, however, the
thetic vasoconstriction is induced (Frey et al. 1980a, b;
diving response serves no purpose and does not set
Allen et al. 1992). These two responses cause an increase
thermoregulatory adjustments aside.
in blood pressure. As the immersion continues, stimu-
lation of the arterial baroreceptors returns the heart rate
Key words Diving response á Bradycardia á
towards the pre-immersion rate, often within the 1st min
Tachycardia á Arterial blood pressure á Cold stimulation
(i.e. the beta-adrenergic sympathetic activity is reduced
while the alpha-adrenergic sympathetic activity contin-
J. Andersson (&) á E. Schagatay á B. Holm ues to result in vasoconstriction and increased blood
Department of Animal Physiology, pressure; Frey et al. 1980a, b).
Lund University, Helgonav. 3B, According to previous studies, chilling of the forearm
22362 Lund, Sweden
e-mail: johan.andersson@zoofys.lu.se and the face has opposite e€ects on heart rate. The
Tel.: +46-46-2229349; Fax: +46-46-2224539 purpose of the study presented here was to investigate
A. GisleÂn
whether forearm immersion in cold water has any e€ect
Department of Zoology, Lund University, on the cardiovascular responses to face immersion and
Lund, Sweden apnoea (the diving response). This was done by

recording the cardiovascular responses to cold-water
immersion of the forearm and face during eupnoea and
apnoea.
Methods
Subjects
A group of 34 healthy subjects, 23 men and 11 women, ranging in
age from 19 to 30 years (mean 23.4 years) participated in this
study. All subjects were non-smokers but two were taking snu€.
Their physical exercise averaged 3.3 h/week, with a range from 0 to
9.5 h/week.
Measurements
An electrocardiogram was checked for anomalies prior to testing
(Cardisuny 501, lead 2, Fukuda ME Kogoyo, Tokyo, Japan). The
subject's vital capacity was measured (Spirolite 201, Vise Medical,
Chiba, Japan), ®rst in the standing position and thereafter with the
subject lying prone on a bed with the equipment attached. Heart
rate and arterial blood pressure were recorded continuously in all
subjects on the left middle ®nger by a photoplethysmometer
(Finapres 2300, Ohmeda, Madison, Wisc., USA). Skin capillary
blood ¯ow in the left thumb was recorded continuously in 22 of the
subjects with a laser Doppler ¯owmeter (Advanced Laser Flow-
meter 21, Advance, Tokyo, Japan). The left hand was positioned at
the same level relative to the heart throughout the experiment
(cf Schagatay and Andersson 1998). Heart rate, blood pressure,
and skin blood ¯ow values were stored continuously in a computer.
Breathing movements were recorded by a pneumatic chest bellows,
which was connected to an ampli®er and an analogue recorder.
Experimental procedures
All experiments were conducted in conformity with the principles
of the Declaration of Helsinki and were approved by the Research
Ethics Committee of the Faculty of Medicine, Lund University.
The experimental protocol was divided into two parts, each part
containing four tests as follows. For part I:
1. Eupnoea with forearm immersion.
2. Eupnoea with face immersion.
3. Eupnoea with forearm and face immersion.
4. Apnoea with face immersion.
And for part II:
1. Apnoea without immersion.
2. Apnoea with forearm immersion.
3. Apnoea with face immersion.
4. Apnoea with forearm and face immersion.
Nineteen subjects performed part I [mean (SD) age 23 (3) years,
height 1.77 (0.1) m, body mass 70 (12) kg, vital capacity 5.3 (1.0) l],
and 23 subjects performed part II [mean (SD) age 24 (2) years,
height 1.79 (0.1) m, body mass 72 (10) kg, vital capacity 5.2 (1.0) l].
Eight subjects participated in both parts of the study. The forearm
and face were immersed in water at 9±11 °C. Room temperature was
22±26 °C. The order in which the tests were conducted was rando-
mised for each subject, and for the eight subjects that participated in
both parts, part I and part II were performed on separate days.
The subjects were familiarised with the equipment and thor-
oughly informed about the experimental procedures, after which an
informed consent form was signed. The subject assumed a prone
position on a mattress with the head resting on a removable pillow
above the face-immersion trough. The subject's arms were resting
on each side of the face-immersion trough. To the right of the bed,
the forearm immersion trough was situated on a small table. If the
567
test did not include forearm immersion, the subject was instructed
to lay his/her arm on the table. During forearm immersion tests,
the right forearm and hand were immersed. If the test did not
include face immersion, the subject was instructed to hold his/her
face just over the water. During face immersions, the entire face
including the forehead and chin were immersed. This assured an
identical position during all the tests. The subject was instructed to
breathe normally throughout the experiment and to make a deep,
but not maximal inspiration when initiating apnoea. In all of the
eupnoea tests the subject started breathing through a snorkel at
least 2 min before the beginning of the test. The individual respi-
ratory frequency and tidal volume during the control period before
the immersions were continued during immersions as judged by
observations of the breathing movements on the analogue recorder.
The recording of all parameters was commenced 2 min prior to
each test. The pillow was removed 15 s prior to the tests. At this
point, the subject was instructed to put his/her right arm in position
beside the trough. A countdown was given during the last 10 s.
During apnoea tests, the initiations of apnoea and immersion were
simultaneous. Immersions without apnoea lasted for 60 s, while
apnoeas were performed to individual maximal duration [mean
(SD) for all apnoeas: 108 (49) s]. After the immersion/apnoea was
terminated, the recording continued for a further 2 min. The pause
between each test comprised 10 min of prone rest.
Data analysis
The control values for heart rate, skin blood ¯ow, and mean
arterial pressure were calculated as time averages from the period
90±30 s before each test. The time averages of these parameters
were also calculated during the period 30±45 s into each test. The
relative changes were calculated between the time averages from the
period 30±45 s into the tests and the corresponding control values.
We chose the period 30±45 s into the tests for comparative analysis
because other studies indicate that the heart rate e€ect of forearm
immersion in cold water is of a short duration (Frey et al. 1980a,
b), and because it takes approximately 30 s for the human diving
response to develop completely (Jung and Stolle 1981; Andersson
and Schagatay 1998b; Schagatay et al. 1999). Thus, the chosen
period was considered as the optimum for investigating the inter-
action between the responses to forearm and face immersion and
their relationship to apnoea. For statistical evaluations within the
two parts of the study, a paired t test with Bonferroni correction for
multiple comparisons was used. For comparisons between the two
apnoeas with face immersions, an unpaired t test was used. The
level of statistical signi®cance was set at P < 0.05.
Results
Development of cardiovascular responses
Part I
During the forearm immersion, the heart rate increased
slightly, but towards the end of the 60-s immersion the
heart rate had returned to the control level (Fig. 1).
The skin capillary blood ¯ow was reduced throughout
the entire forearm immersion. The mean arterial pressure
increased during the forearm immersion. During the face
immersion while breathing through a snorkel, a similar
pattern of cardiovascular responses could be observed,
except for the heart rate response. After an initial
tachycardic response, a heart rate reduction well below
the control level developed. During the concurrent face
and forearm immersion, the skin blood ¯ow and blood
568
pressure followed the same pattern as during forearm or
face immersion, while the heart rate response was of an
intermediate magnitude compared to during forearm or
face immersion. After an initial tachycardic response,
apnoea with face immersion induced a de®nite heart rate
reduction. The skin blood ¯ow during apnoea with face
immersion followed the same pattern as during the other
tests, while the increase in blood pressure was more
marked during the apnoea than in any other test in part I.
Part II
During all of the apnoeas, a heart rate reduction followed
an initial tachycardic response that started even before
the beginning of apnoea (Fig. 2). The heart rate reduc-
Fig. 1 Mean heart rate, skin capillary blood ¯ow, and mean arterial
pressure before, during, and after tests I1±I4: I1 eupnoea with forearm
immersion; I2 eupnoea with face immersion; I3 eupnoea with face and
forearm immersion; I4 apnoea with face immersion. Heart rate and
arterial pressure: n ˆ 19; Skin blood ¯ow: n ˆ 14. Downward-pointing
arrows indicate the start of the tests, and upward-pointing arrows
indicate the end of the tests. The break in the line of test I4 indicates
that the apnoea was of a longer duration than 60 s
tions observed during the apnoeas without face immer-
sions were not as marked as those observed during the
apnoeas with face immersions. The skin capillary blood
¯ow was reduced during all of the apnoeas. During the
apnoea without immersion, the ®nal level of skin blood
¯ow adjustment was higher than in the other tests. After
initial reductions during the ®rst 20 s, the mean arterial
pressure increased throughout the apnoeic episodes.
Again, the responses during apnoeas without face im-
mersions were not as marked as in the other two tests.
Anticipatory responses
For both heart rate and skin blood ¯ow, anticipatory
responses beginning about 30 s prior to apnoeas or
Fig. 2 Mean heart rate, skin capillary blood ¯ow, and mean arterial
pressure before, during, and after tests II1±II4: II1 apnoea without
immersion; II2 apnoea with forearm immersion; II3 apnoea with face
immersion; II4 apnoea with forearm and face immersion. Heart rate
and arterial pressure: n ˆ 23; Skin blood ¯ow: n ˆ 16. Downward-
pointing arrows indicate start of the tests, and upward-pointing arrows
indicate the end of tests. The break in the lines indicates that the
apnoeas were of longer duration than 60 s
 569

immersions could be observed. We performed additional
experiments on six subjects to elucidate the e€ects of
``dummy immersions'' on the anticipatory responses.
With the same experimental set-up, after an identical
countdown, subjects had the forearm lowered (by the
experimenter) into a trough with or without cold water.
Subjects were unaware of the presence or absence of the
water until the forearm was lowered into the trough, as
con®rmed by post-test interviews. Results similar to
those observed in the main experiment were observed
during the ``true immersion'' (Fig. 3). The heart rate
increased slightly, by 1%, compared to the control level
during immersions, and the skin blood ¯ow was reduced
by 54%. During the ``dummy immersion'', the cardio-
vascular parameters followed a di€erent pattern (Fig. 3).
The heart rate decreased by 4% and the skin blood ¯ow
increased by 15%.
forearm immersion. Within part II, the apnoeas without
face immersions resulted in less pronounced heart rate
reductions when compared to the apnoeas with face
immersions (P < 0.001). However, heart rate responses
during the apnoea in air and the apnoea with forearm
immersion did not di€er. Neither did the responses
during the apnoea with face immersion and the apnoea
with face and forearm immersion di€er.
Skin blood ¯ow
The relative changes in skin capillary blood ¯ow did not
di€er among tests I1±4 (Fig. 4). However, the skin blood
¯ow reduction during the apnoea in air was less
pronounced than the reductions during the apnoea with
forearm immersion and apnoea with face and forearm
immersion (P < 0.05).

Comparisons between cardiovascular responses

to immersion and apnoea

There were no di€erences in the cardiovascular

responses to apnoea with face immersion between parts I
and II.

Heart rate

The relative changes in heart rate from the control levels

during the period 30±45 s into the tests were compared
(Fig. 4). Within part I, the apnoea with face immersion
resulted in a heart rate response that di€ered from all the
other tests (P < 0.01). The forearm immersion and the
face immersion resulted in di€erent responses
(P < 0.01). The responses in these two tests did not
di€er from the response during concurrent face and

Fig. 3 Mean skin capillary blood ¯ow before, during, and after the
forearm was lowered into a trough with (true immersion, solid line) or
without cold water (dummy immersion, dashed line). Values are means
from six subjects. Downward-pointing arrows indicate start of the
immersions, and upward-pointing arrows indicate the end of immer-
sions.
Fig. 4 Percentage changes in heart rate [n ˆ 19 (part I) or n ˆ 23
(part II)], skin capillary blood ¯ow [n ˆ 14 (part I) or n ˆ 16 (part II)],
and mean arterial pressure [n ˆ 19 (part I) or 23 (part II)] from the
control period for tests I1±I4 and II1±II4 (see Figs. 1, 2). Data are
presented as the means (SEM). *P < 0.05, **P < 0.01,
***P < 0.001
570
Mean arterial pressure
The mean arterial pressure increased more (P < 0.01)
during the apnoea than in any other test within part I
(Fig. 4). During the apnoea with face and forearm im-
mersion, the mean arterial pressure increased more than
in the apnoeas without face immersions (P < 0.05).
Discussion
Earlier studies have shown that the cardiovascular
responses to cold stimulation of the extremities can be
additive to the responses from other cardiovascular
challenges (e.g. changes in central blood volume and
isometric exercise; Ebert et al. 1982; Peikert and Smo-
lander 1991). There has been, to our knowledge, no
other study investigating whether cold stimulation of the
extremities a€ects the cardiovascular responses to
apnoea, the human diving response. Our results indicate
that during eupnoea, the heart rate response to simul-
taneous cold stimulation of the forearm and face is an
additive response to the two stimuli, while the skin
blood ¯ow and arterial pressure responses are not fur-
ther augmented by the combined stimulation. During
apnoea the pattern changed. Instead of an additive re-
sponse, a complete development of the human diving
response occured (i.e. cold stimulation of the forearm
does not attenuate the heart rate reduction, nor does it
augment the skin blood ¯ow or blood pressure re-
sponse).
Cold stimulation of the extremities elicits ther-
moregulatory re¯exes that result in a reduction in heat
loss to the environment. One of the most prominent
responses is a reduction in peripheral blood ¯ow. As the
result of a general sympathetic activation, the heart rate
and heart contractility is increased (Frey et al. 1980a).
The vasoconstriction and increased cardiac output leads
to an increase in the arterial blood pressure. This pattern
of cardiovascular re¯exes could be observed in the
results of this study. Immersion of the face in cold water
elicited a di€erent cardiovascular response. Stimulation
of the facial cold receptors resulted in a heart rate
reduction and a reduced peripheral blood ¯ow. These
autonomic re¯exes are brought about by vagal chrono-
tropic e€ects on the heart and alpha-adrenergic sympa-
thetic out¯ow to resistance vessels (Finley et al. 1979;
Frey et al. 1980a; Fagius and Sundlof 1986). Since the
reduced cardiac output in this situation was not
balanced to the increased total peripheral resistance, the
arterial blood pressure was elevated. When the two
stimuli (forearm and face immersion) were combined, an
additive response pattern was observed regarding the
heart rate. The heart rate during concurrent forearm and
face immersion was at an intermediate level compared to
during the forearm or face immersion. However, the
vasoconstriction obtained during either the forearm or
face immersion was not further augmented by cold
stimulation of both areas simultaneously, as indicated
by the lack of di€erences in skin capillary blood ¯ow
among the tests of part I in this study (Figs. 1, 4).
The heart rate reduction and selective vasoconstric-
tion elicited by apnoea without immersion were aug-
mented by immersion of the face in cold water, as
reported previously (Kawakami et al. 1967; Hurwitz and
Furedy 1986; Marsh et al. 1995; Andersson and Sch-
agatay 1998a). However, forearm immersion did not
a€ect the cardiovascular responses during apnoea as it
did during eupnoea (i.e. the human diving response was
una€ected by cold stimulation of the forearm during the
apnoeas). When comparing the response to apnoea with
that obtained during the apnoea with forearm immer-
sion, we found no di€erence in heart rate or blood
pressure responses. Neither were any di€erences found
when the responses during the apnoea with face
immersion and the apnoea with face and forearm
immersion were compared. Therefore, we conclude that
forearm immersion has no e€ect on the development of
the general diving response when immersed simulta-
neously as apnoea begins.
The reduction in skin capillary blood ¯ow during
apnoea was augmented by cold-water immersion. In
fact, all immersions resulted in similar skin capillary
blood ¯ow levels, during both eupnoea and apnoea.
Cold stimulation is the most e€ective stimulus causing
re¯ex vasoconstriction in the skin (Heistad and Abboud
1974). Thus, our results indicate that cold stimulation,
regardless of the site of stimulation, elicits a vasocon-
striction in skin vessels that can not be further aug-
mented by simultaneous cold stimulation of another
part of the body surface.
The mean arterial blood pressure was more elevated
during the apnoeas than during the immersions with
eupnoea. It has been shown that the cardiac output is
reduced during breath holding (Ferrigno et al. 1986).
The increased blood pressure thus implies that the total
peripheral resistance has increased more than the
cardiac output decreased. Thermal stimuli primarily
a€ect skin blood ¯ow (Heistad and Abboud 1974).
Apnoea also a€ects blood ¯ow in other vascular beds
(e.g. muscle blood ¯ow; Zbrozyna and Westwood
1992). The ®ndings of higher arterial pressures during
the apnoeas compared to the eupnoeic immersions
could thus be due to a vasoconstriction in vessels not
a€ected by thermal stimuli. At the onset of apnoea,
there was a transient reduction in blood pressure. This
is caused by the elevated intrathoracic pressure that
follows holding the breath with a large lung volume,
initially impeding the venous return and consequently
reducing stroke volume and cardiac output (LineÂr 1994;
Andersson and Schagatay 1998b).
There were anticipatory responses during the count-
down before the tests, but true physiological responses
developed during the immersions and apnoeas. Antici-
patory responses were also clearly visible before the
``dummy immersion'', in particular as transient reduc-
tions in skin blood ¯ow after the instructions ``60 s left''
and ``30 s left'' (Fig. 3). However, when the subjects

Fig. 5 Schematic model for the integration of the cardiovascular
re¯exes evoked by the three stimuli (cold stimulation of the forearm,
cold stimulation of the face, and apnoea). (i Inhibition, + increase,
) decrease)
were surprised by the absence of water, the cardiovas-
cular parameters promptly returned to the pre-count-
down level. In particular, the skin blood ¯ow, which also
showed the most marked anticipatory responses, in-
creased almost instantaneously towards the control level
during the dummy immersion.
Based on the results from this and previous studies, a
schematic model for the integration of the cardiovascular
re¯exes elicited by the three stimuli (cold stimulation of
the forearm, cold stimulation of the face, and apnoea) is
suggested (Fig. 5). During eupnoea the heart rate reduc-
tion derived from chilling of the face is attenuated if the
forearm is chilled simultaneously. Thus, the increased
chronotropic sympathetic stimulation evoked by arm
immersion counteracts the chronotropic parasympathetic
activity induced by facial chilling. The peripheral blood
¯ow reduction is not further augmented by cold stimula-
tion of yet another skin area (i.e. the maximal reduction in
skin capillary blood ¯ow during immersion in cold water
is obtained by either immersion of the face or forearm
separately). During apnoea, on the contrary, the forearm
immersion has no e€ect on the heart rate response. Thus, a
heart rate reduction was induced by an increased para-
sympathetic activity, whereas the sympathetic stimulation
evoked by forearm immersion was not expressed. The
augmented increase in blood pressure during apnoea
compared to that during eupnoeic immersions might be
explained by a vasoconstriction in speci®c vascular beds,
571
besides the reduction in skin blood ¯ow, which would
further increase the total peripheral resistance.
The main conclusions from this study are that during
eupnoea, heart rate responses from forearm and face
immersions are additive, while the heart rate responses
to apnoea are not attenuated by the forearm immersion.
The fact that the oxygen-conserving diving response has
priority over thermoregulatory responses in the threat of
asphyxia appears to be a functional property. During
breathing, however, the diving response serves no pur-
pose, since oxygen delivery is undisturbed and ther-
moregulatory adjustments will be more appropriate
during cold stimulation.
Acknowledgements This study was supported by grants from the
Hierta-Retzius' Foundation for Scienti®c Research, the Royal
Physiographic Society, and the Swedish National Centre for Re-
search in Sports. All experiments comply with the current laws of
Sweden.
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